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1.
Perhaps the most common form of cooperation among primates is the formation of coalitions. Competition among males within a group concerns a constant quantity of the limiting resource (fertilizations). Contest competition over fertilizations is known to produce payoffs that are distributed according to the priority-of-access model, and hence show an exponential decline in payoff with rank. We develop a model for rank-changing, within-group coalitions among primate males. For these coalitions to occur, they must be both profitable (i.e. improve fitness) for all coalition members and feasible (i.e. be able to beat the targets). We assume that the value of the coalition is the sum of the payoffs of the partners in their original ranks. We distinguish three basic coalition configurations, depending on the dominance ranks of the coalition partners relative to their target. We predict five basic coalition types. First, all-up, rank-changing coalitions targeting individuals ranking above all coalition partners; these are expected to involve coalition partners ranking just below their target, concern top rank, and be small, just two or three animals. Second, bridging, rank-changing coalitions, where higher-rankers support lower-rankers to rise to a rank below themselves; these are expected to be most common where a high-ranking male in a despotic system can support a low-ranking relative. Third, bridging non-rank-changing coalitions; these are expected to be common whenever high-ranking males have low-ranking close relatives. Fourth, non-rank-changing coalitions by high-rankers against lower-ranking targets; these are expected to serve to counteract or prevent the first type. Fifth, non-rank-changing, leveling coalitions, in which all partners rank below their target and which flatten the payoff distribution; these are expected to be large and mainly involve lower-ranking males. Bridging, rank-changing coalitions are expected in situations where contest is strong, all-up rank-changing coalitions where contest is intermediate, and leveling coalitions where contest is weak. We review the empirical patterns found among primates. The strong predictions of the model are confirmed by observational data on male-male coalitions in primates.
Carel P. van SchaikEmail:
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2.
As applied to polygynous mammals, the socioecological model assumes that environmental risks and resources determine the spatial and temporal distribution of females, which then sets male strategies for monopolizing fertile matings. The effects of female spatial distribution (i.e., female number) and temporal overlap (female mating synchrony) have been examined in comparative studies of primates, but the relative influence of these two factors on male monopolization potential (the number of males) remains unclear. One particular problem is that female synchrony is more difficult to estimate than female number. This paper uses multivariate statistical methods and three independent estimates of female synchrony to assess the roles of spatial and temporal effects in the context of a phylogenetically corrected dataset. These analyses are based on sensitivity analyses involving a total of four phylogenies, with two sets of branch length estimates for each tree, and one nonphylogenetic analysis in which species values are used (because male behavior may represent a facultative response to the distribution of females). The results show: (1) that breeding seasonality predicts male number (statistically significant in six out of nine sensitivity tests); (2) that expected female overlap, after controlling for female group size using residuals, also accounts for the number of males in primate groups (significant in eight out of nine tests), and (3) that actual estimates of female mating synchrony predict male number, again after correcting for female group size (significant in five out of nine tests). Nonsignificant results are in the predicted direction, and female group size is significant in all statistical tests. These analyses therefore demonstrate an independent influence of female temporal overlap on male monopolization strategies in mammalian social systems. Received: 24 July 1998 / Received in revised form: 5 February 1999 / Accepted: 7 February 1999  相似文献   

3.
In order to remain stable, dispersed social groups have to solve two fundamental problems: the coordination of movement and cohesiveness within a group and the spacing between the groups. Here, we investigate mechanisms involved in intra-group coordination and inter-group spacing using the golden brown mouse lemur, Microcebus ravelobensis, as a model for a nocturnal, solitary foraging mammal with a dispersed social system. By means of radiotelemetry and bioacoustics we studied the olfactory and vocal behaviour during nocturnal dispersal and reunion of five sleeping groups.All groups used 3–17 sleeping sites exclusively, suggesting a sleeping site-related territoriality and competition for them. The occurrence of olfactory and vocal behaviour showed an asymmetrical temporal distribution. Whereas marking behaviour was observed exclusively during dispersal, a particular call type, the trill, was used by all groups during reunions. Interestingly, these trills carried group-specific signatures.Our findings provide the first empirical evidence for nocturnal primates in a natural environment that olfactory signals represent an important mechanism to regulate the distribution of different groups in space, whereas acoustic signals control intra-group cohesion and coordination.  相似文献   

4.
The number of adult males in a bisexual primate group is thought to be determined mainly by the defensibility of mating access to the females. However, among primates, small groups sometimes contain several adult male. We evaluate the hypothesis that high predation risk may lead to greater male representation in primate groups, independent of the effect of group size. Male primates are known to be more vigilant than females, and may also be better at detecting potential predators. A comparative test was done using arboreal folivores in three continents, howler monkeys in the Neotropics, colobus monkeys in Africa and langurs in South and Southeast Asia. Howlers and colobus are exposed to predation by large monkey-eating eagles, whereas the langurs are not. We conducted regression analyses on group composition data of single populations and on mean group compositions among populations, and found that the arboreal langurs basically live in groups with only a single adult males whereas groups of howler and colobus monkeys of comparable size have multiple adult males (Fig. 3). Thus, the hypothesis was supported. Several alternative hypotheses do not account for this pattern.  相似文献   

5.
The population dynamics of species interactions provides valuable information for life sciences. Lotka-Volterra equations (LVEs) are known to be the most popular model, and they are mainly applied to the systems of predation and competition. However, LVEs often fail to catch the population dynamics of mutualism; the population sizes of species increase infinitely under certain condition (divergence problem). Furthermore, LVEs never predicts the Allee effect in the systems of obligate mutualism. Instead of LVEs, several models have been presented for mutualism; unfortunately, they are rather complicated. It is, therefore, necessary to introduce a simpler theory for mutualism. In the present paper, we apply the lattice gas model which corresponds to the mean-field theory of the usual lattice model. The derived equations are cubic and contain only essential features for mutualism. In the case of obligate mutualism, the dynamics exhibits the Allee effect, and it is almost the same as in the male-female systems. In our model, the population sizes never increase infinitely, because our model contains not only intra- but also interspecific competitions. If the density of one species increases disproportionately in respect of its mutual partners, then this might imply downward pressure on the population abundance of the mutual partner species and such feedback would eventually act as a controlling influence on the population abundance of either species. We discuss several assumptions in our model; in particular, if both species can occupy in each cell simultaneously, then the interspecific competition disappears.  相似文献   

6.
In temperate climates groundwater can have a profound effect on vegetation, because it strongly influences the spatio-temporal distribution of soil moisture in the rootzone and therefore the occurrence of water and oxygen stress of vegetation. This article focuses on vegetation and groundwater dynamics along a hill slope by developing and evaluating a fully coupled hydrological-vegetation model for a temperate forest ecosystem. The vegetation model is described in part 1 of this series of two papers. To simulate the hydrology an extended version of the saturated-unsaturated hydrological model STARWARS has been used. The coupled model is used to investigate both the short and long-term dynamics for a system of two species. Both compete for light and water where one is adapted to wet conditions and the other to dry conditions. The daily dynamics show that the influence of groundwater is particularly strong in spring when waterlogging occurs due to decreased evapotranspiration in winter. Long simulation runs of 1000 years were performed to study the equilibrium state for the two species. Comparison of simulation results with observations of groundwater depth and vegetation types along a dry-wet gradient in a natural forest shows that a reductionist approach is able to capture these patterns well. Sensitivity analysis shows that the border between wet- and dry-adapted species moves upslope with increased rainfall, decreased slope angle and decreased aquifer thickness. These results are similar to previous findings which were based on global maximization of ecosystem evaporation or minimizing ecosystem stress. Comparison of runs with a fixed and a dynamic groundwater table shows that a dynamic groundwater table facilitates a wider transition zone between vegetation types along the hill slope. In this transition the biomass of vegetation is higher in the case of a dynamic groundwater than in case of a static groundwater table. This underlines the importance of incorporating spatial groundwater dynamics in models of groundwater influenced ecosystems.  相似文献   

7.
Vegetation growth models often concentrate on the interaction of vegetation with soil moisture but usually omit the influence of groundwater. However the proximity of groundwater can have a profound effect on vegetation growth, because it strongly influences the spatial and temporal distribution of soil moisture and therefore water and oxygen stress of vegetation. In two papers we describe the behavior of a coupled vegetation-groundwater-soil water model including the competition for water and light. In this first paper we describe the vegetation model, compare the model to measured flux data and show the influence of water and light competition in one dimension. In the second paper we focus on the influence of lateral groundwater flow and spatial patterns along a hillslope. The vegetation model is based on a biophysical representation of the soil-plant-atmosphere continuum. Transpiration and stomatal conductance depend both on atmospheric forcing and soil moisture content. Carbon assimilation depends on environmental conditions, stomatal conductance and biochemical processes. Light competition is driven by tree height and water competition is driven by root water uptake and its water and oxygen stress reaction. The modeled and measured H2O and CO2 fluxes compare well to observations on both a diurnal and a yearly timescale. Using an upscaling procedure long simulation runs were performed. These show the importance of light competition in temperate forests: once a tree is established under slightly unfavorable soil moisture conditions it can not be outcompeted by smaller trees with better soil moisture uptake capabilities, both in dry as in wet conditions. Performing the long simulation runs with a background mortality rate reproduces realistic densities of wet and dry adapted tree species along a wet to dry gradient. These simulations show that the influence of groundwater is apparent for a large range of groundwater depths, by both capillary rise and water logging. They also show that species composition and biomass have a larger influence on the water balance in eco-hydrological systems than soil and groundwater alone.  相似文献   

8.
基于盲数的水体沉积物潜在生态风险评价方法   总被引:2,自引:0,他引:2  
李如忠  洪天求  贾志海  刘路 《生态环境》2007,16(5):1346-1352
基于生态风险评价系统中多种不确定性共存或交叉存在的特性,将盲数理论应用于水体沉积物潜在生态风险的评价研究。在将Hakanson潜在生态风险指数模型参数定义为盲参数的基础上,建立了沉积物潜在生态风险评价的盲数模型,并提出了污染程度和潜在生态风险等级识别盲数可靠性计算模型。根据上述盲数模型,不仅可以得到沉积物污染程度和潜在生态风险的可能取值区间,也可以得到各区间相应的可信度水平,进而定量计算待评价对象隶属于各评价等级的主观可能性大小。作为案例,将上述模型应用于巢湖十五里河河口沉积物中重金属潜在生态风险评价的研究。在得到沉积物中单个重金属污染物(Hg,Cd,Pb,Cu,Cr)隶属于污染程度各等级和潜在生态风险各等级可信度水平的基础上,得出十五里河河口处于较高生态风险水平的结论,相应的可信度为0.626。实例研究表明,以盲数理论评价沉积物的潜在生态风险,理论上可行,评价结论可信。  相似文献   

9.
This article presents a system dynamics (SD) method to examine the problem of forest degradation. The model developed takes a system-oriented view of forest management, embracing both social and biophysical factors affecting deforestation. Social factors examined are socio-economic variables or elements that influence behaviour and decision-making choices at the household level. Biophysical factors are four sub-components that are considered major land uses namely, the paddy field component, rattan plantations, coffee plantations and forest stands. The model was applied in a case study located in Pasir District of East Kalimantan, Indonesia. The site covers an area that includes a protected forest and a privately allocated timber license concession. Three village communities are examined in the case study. The SD model developed was applied to the case study focusing on three management policies or scenarios, which are based on access rights to the forest resources within the study area. Specifically, the property arrangements examined in each scenario are: Policy 1 – status quo (i.e. continue present property rights arrangements); Policy 2 – local communities manage the forest exclusively; and Policy 3 – collaborative management involving both local communities and a private company. Results from the model show that the third policy is the most viable option, and also lead to a win–win solution.  相似文献   

10.
Group sizes are often considered to be the result of a trade-off between predation risk and the costs of feeding competition. We develop a model to explore the interaction between different ecological constraints on group sizes, using a primate (baboons) case study. The model uses climatic correlates of time budgets to predict maximum ecologically tolerable group size, and climatic predictors of predation risk (reflected mainly in predator density and female body mass) to predict minimum tolerable group size for any given habitat. As well as defining the range of sustainable group sizes for a given habitat, the model also allows us to reliably predict our exemplar taxon's biogeographical distribution across Africa. We also explore the life history implications of the model to ask whether baboons form group sizes which maximise survival or fecundity in the classic trade off between these two key life history variables. Our results indicate that, within the range of study sites in our sample, baboons prefer to maximise fecundity. However, the data indicate that in higher predation risk habitats they would switch to maximising survival at the expense of fecundity. We argue that this is due to the fact that interbirth interval and developmental rates have a ceiling that cannot be breached. Thus, while females can shorten interbirth intervals to compensate for increased predation risk, there is a limit to how much these life history variables can be altered, and when this is reached the best strategy is to maximise survivorship.  相似文献   

11.
The models used for ecosystems modeling are generally based on differential equations. However, in recent years new computational models based on biological processes, or bioinspired models, have arisen, among which are P systems. These are inspired by the functions of cells and present important advantages with respect to traditional models, such as a high computational efficiency, modularity and their ability to work in parallel. They are simple, individual-based models that use biological parameters that can be obtained experimentally. In this work, we present the framework for a model based on P systems applied to the study of an ecosystem in which three avian scavengers (predators) interact with 10 wild and domestic ungulates (preys). The computation time for 100 repetitions, corresponding to 14 simulation years each, with an initial population composed of 385,422 individuals, was 30 min. Our results suggest that the model presented, based on P systems, correctly simulates the population dynamics in the period of time analyzed. We discuss the usefulness of this tool in simulating complex ecosystems dynamics to aid managers, conservationists and policy-makers in making appropriate decisions for the improvement of management and conservation programs.  相似文献   

12.
非稳态条件下藻类种间非生物资源竞争理论及研究进展   总被引:2,自引:0,他引:2  
金相灿  杨苏文  姜霞 《生态环境》2007,16(2):632-638
从理论及实验研究的角度论述了藻类非稳态种间资源竞争研究进展。实验室内稳态条件下的竞争理论得到了大量实验验证,由于自然水体处于非稳状态,非稳态条件下藻类种间资源竞争日益成为藻类水华形成机制研究的重点。主要理论包括Tilman的R*法则和资源比假说,Grover的拾遗者-机会主义者交替竞争假说(VIS),中度干扰假说(IDH)及关键光强(CLIH)假说。这些假说的提出丰富了藻类资源竞争的理论体系,为研究自然状态下藻类增殖和竞争行为奠定理论基础。今后的藻类种间资源竞争的研究重点为非稳态下物理条件变化以及物理、化学和生物资源共同变化下藻类种间关系及生物多样性的研究。  相似文献   

13.
Several reaction schemes, based on the conserved scalar theory, are implemented within a stochastic Lagrangian micromixing model to simulate the dispersion of reactive scalars in turbulent flows. In particular, the formulation of the reaction-dominated limit (RDL) reaction scheme is here extended to improve the model performance under non-homogeneous conditions (NHRDL scheme). The validation of the stochastic model is obtained by comparison with the available measurements of reactive pollutant concentrations in a grid-generated turbulent flow. This test case describes the dispersion of two atmospheric reactant species (NO and O3) and their reaction product (NO2) in an unbounded turbulent flow. Model inter-comparisons are also assessed, by considering the results of state-of-the-art models for pollutant dispersion. The present validation shows that RDL reaction scheme provides a systematic overestimation (relative error of ca. 85% around the centreline) in computing the local reactant consumption/production rate, whereas the NHRDL scheme drastically reduces this gap (relative error lower than 5% around the centreline). In terms of NO2 production (or reactant consumption), neglecting concentration fluctuations determines overestimations of the product mean of around 100% and a NO2 local production of one order of magnitude higher than the reference simulation. In terms of standard deviations, the concentration fluctuations of both the passive and reactive scalars are generally of the same order of magnitude or up to 1 or 2 orders of magnitudes higher than the corresponding ensemble mean values, except for the background reactant close to the plume edges. The study highlights the importance of modelling pollutant reactions depending on the instantaneous instead of the mean concentrations of the reactants, thus quantifying the role of the turbulent fluctuations of concentration, in terms of scalar statistics (mean, standard deviation, intensity of fluctuations, skewness and kurtosis of concentration, segregation coefficient, simulated reaction rate). This stochastic particle method represents an efficient numerical technique to solve the convection–diffusion equation for reactive scalars and involves several application fields: micro-scale air quality (urban and street-canyon scales), accidental releases, impact of odours, water quality and fluid flow industrial processes (e.g. combustion).  相似文献   

14.
The red bishop (Euplectes orix) is a highly polygynous and colonial weaverbird. Males construct several nests within their territories to which they try to attract females, and females are solely responsible for incubation and raising offspring. In this paper, we describe the characteristics of the red bishop’s mating system as a biological market and investigate the role of nests built by males as a traded commodity in a mating market. As timing of breeding in red bishops in arid and semi-arid zones depends on rainfall patterns which are often unpredictable, there are temporal changes in demand for and supply of nests within a breeding season, with breeding activities of males and females being highly synchronised. We found that males increased their nest-building speed with increased female breeding activity independently of rainfall, indicating that supply follows demand in this mating market. The supply of nests was always larger than the demand for nests. Construction costs for nests increased with demand for nests as indicated by shorter nest-building duration and shorter building delays between two consecutively built nests at times of high breeding activity. Males as a trading class are chosen according to the age of their nests offered, with young nests having a higher probability of being accepted by females. Furthermore, female choosiness with regard to nest age decreased when their own market value decreased, as predicted by biological market theory. The temporal changes of breeding activity together with the female preference for young and fresh nests require that males quickly adjust nest-building activity to varying female demand for new nests. However, males with a better adjustment of building speed to female breeding activity did not gain higher mating success.  相似文献   

15.
Information on benthic carbon mineralization rates is often derived from the analysis of oxygen microprofiles in sediments. To enable a direct comparison of different sediment environments, it is often desirable to characterize sediments by a single proxy that expresses their “reactivity” towards oxygen. For this, there are three commonly used proxies: the oxygen penetration depth (OPD), the oxygen flux at the sediment-water interface (DOU), and the maximum volumetric oxygen consumption rate (Rmax). The OPD can be directly determined from the oxygen depth profile, while the DOU is usually obtained by a linear fit to the oxygen gradient either in diffusive boundary layer. The oxygen consumption rate Rmax requires the fitting of a reactive-transport model to the data profile. This article shows that the OPD alone is a suboptimal proxy, because it shows a strong dependence on the half-saturation constant Ks, and secondly, because it is sensitive to the particular re-oxidation conditions right above the oxic-anoxic interface. Similarly, the volumetric oxygen consumption rate Rmax is rather strongly dependent on the kinetic model formulation employed. To show this we fitted three different (Bouldin, Blackman and Monod) kinetics to the same oxygen data profiles. When fitting these models, the Rmax values obtained differed by 20% for exactly the same oxygen profile. Accordingly, if one reports Rmax values, it is crucial to specify the kinetic model alongside. Overall, DOU emerges as sediment reactivity proxy which is the least model dependent.  相似文献   

16.
The self-thinning line is a very robust pattern, which can be obtained in modeling studies by a variety of different mechanistic assumptions. Our opinion is that we can only advance in our understanding of mechanisms leading to the self-thinning relationship if we demand that the model also reproduces several other characteristic features (patterns) of the self-thinning process such as the degree of size inequality and the average size. We use a pattern-oriented modeling approach to develop a model of self-thinning under size inequality in overcrowded, even-aged stands, which reproduces these three patterns simultaneously. Our approach is to first develop an initial model based on our current ecological knowledge and then to refine the model by modifying the initial model to derive the model that reproduces all patterns of interest.The initial model is as simple as possible while avoiding incidental, ecologically unjustified, assumptions. It is a further development of zone of influence-simulation models: each plant is described by two circles, one describing a minimum-domain-area and one describing the zone of influence. In the initial model, mortality is “death-by-contact” of minimum-domain-areas and growth is a function of inter-tree competition, i.e. overlapping zones of influence. Model parameterization is based on field data on Acacia reficiens in southern Africa. Simulations follow patches of initially small trees through time for up to 1000 years with five parameters, three describing growth and two describing inter-tree competition. A sensitivity analysis shows that all parameters of the initial model contribute significantly to the number and size of plants through time. The two competition parameters, which describe competitive asymmetry and the size of the zone of influence relative to canopy size, are both important for generating size inequality. Thus, both competitive asymmetry and spatial pattern contribute to size inequality, and their relative importance may vary greatly.The sensitivity analysis suggests that all processes included in the initial model are essential to the evolution of size inequality. However, size inequality under the initial model is below field values, meaning that additional, as yet unconsidered processes, contribute to size inequality. Our best-fit model additionally contains details on growth stochasticity.This study establishes the often-proposed direct link between mortality driven by local competition and self-thinning and highlights the importance of stochasticity in ecological processes.  相似文献   

17.
Global positioning system (GPS) collars have revolutionized the collection of animal location data; however, it is well-recognized that considerable bias can be present in these data due to habitat or behavior-induced obstruction of satellite signals resulting in inaccurate or missing locations. To date, no explicit theoretical framework of GPS fix acquisition specific to animal telemetry has been presented, and studies make differing assumptions regarding factors influencing GPS fix acquisition and how these data should be analyzed. Inappropriate statistical models have been used, interaction effects have been misunderstood, and the implementation of bias mitigation techniques has been problematic. Herein we outline current conceptual and analytical problems in the GPS animal telemetry literature, and subsequently present a theoretical model-based framework for GPS fix acquisition that clarifies the single and interactive effects of habitat and behavioral obstruction, fix interval, and collar model on GPS collar performance. By recognizing that GPS fix acquisition is a Bernoulli process, it becomes apparent that all forms of obstruction inherently interact with each other, making generalizations across study areas, study species, and collar models problematic. Stationary collar tests to determine the probability of fix acquisition (PFA), location accuracy, and the response to sources of obstruction are thus of limited applicability to animal-deployed collars. Bias mitigation techniques that extrapolate PFA models across samples, especially those using stationary collar tests to correct animal-deployed collars, are theoretically unsound. It is also demonstrated that nonlinearities in the relationships between sources of obstruction and PFA complicate PFA modeling with limited data and that even slight model misspecification can lead to considerable errors in correction factors, especially when using inverse weighting to mitigate bias. By emphasizing the importance of GPS collar sensitivity and ephemeris retention, the theoretical framework predicts that newer, more sensitive GPS collars will be less severely biased by sources of obstruction than reported for the older, less sensitive collars that have been used in the majority of GPS performance studies to date and we expect this trend to continue. This heuristic modeling exercise should be of value to researchers planning and analyzing studies using GPS collars and it also establishes a starting point for future theoretical investigations into GPS collar performance and bias mitigation.  相似文献   

18.
19.
In this study we combined an extensive database of observed wildfires with high-resolution meteorological data to build a novel spatially and temporally varying survival model to analyze fire regimes in the Mediterranean ecosystem in the Cape Floristic Region (CFR) of South Africa during the period 1980-2000. The model revealed an important influence of seasonally anomalous weather on fire probability, with increased probability of fire in seasons that are warmer and drier than average. In addition to these local-scale influences, the Antarctic Ocean Oscillation (AAO) was identified as an important large-scale influence or teleconnection to global circulation patterns. Fire probability increased in seasons during positive AAO phases, when the subtropical jet moves northward and low level moisture transport decreases. These results confirm that fire occurrence in the CFR is strongly affected by climatic variability at both local and global scales, and thus likely to respond sensitively to future climate change. Comparison of the modelled fire probability between two periods (1951-1975 and 1976-2000) revealed a 4-year decrease in an average fire return time. If, as currently forecasted, climate change in the region continues to produce higher temperatures, more frequent heat waves, and/or lower rainfall, our model thus indicates that fire frequency is likely to increase substantially. The regional implications of shorter fire return times include shifting community structure and composition, favoring species that tolerate more frequent fires.  相似文献   

20.
Recent calls for the development of ecosystem-based fisheries management compel the development of resource management tools and linkages between existing fisheries management tools and other resource tools to enable assessment and management of multiple impacts on fisheries resources. In this paper, we describe the use of the Chesapeake Bay Fisheries Ecosystem Model (CBFEM), developed using the Ecopath with Ecosim (EwE) software, and the Chesapeake Bay Water Quality Model (WQM) to demonstrate how linkages between available modeling tools can be used to inform ecosystem-based natural resource management. The CBFEM was developed to provide strategic ecosystem information in support of fisheries management. The WQM was developed to assess impacts on water quality. The CBFEM was indirectly coupled with the WQM to assess the effects of water quality and submerged aquatic vegetation (SAV) on blue crabs. The output from two WQM scenarios (1985-1994), a baseline scenario representing actual nutrient inputs and another with reduced inputs based on a tributary management strategy, was incorporated into the CBFEM. The results suggested that blue crab biomass could be enhanced under management strategies (reduced nutrient input) when the effective search rate of blue crab young-of-the-year's (YOY's) predators or the vulnerability of blue crab YOY to its predators was adjusted by SAV. Such model linkages are important for incorporating physical and biological components of ecosystems in order to explore ecosystem-based fisheries management options.  相似文献   

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